"Whoever wants to know the age at which newborn cheetah kittens open their eyes, the size of tiger territories and dispersal patterns of pumas, the food habits of pampas cats, the scent-marking methods of ocelots, and any number of other facts will find them in this book. It is a valuable compendium testifying to the diligence of the authors."George B. Schaller, Wildlife Conservation Society

"This encyclopedic work summarizes all current knowledge about the ecology, behavior, and reproduction of every wild felid species in the world. The authors are eminently qualified and the science throughout the book is excellent; they have done a Herculean job of accurately distilling an enormous literature in a manner that is equally accessible to both a specialist or the curious owner of a housecat."Louise Emmons, Smithsonian National Museum of Natural History

An excerpt fromWild Cats of the Worldby Mel Sunquist and Fiona Sunquist

The Domestic CatHistory, Folklore, Ecology, and Behavior

Over the course of recent history, humans have tamed and domesticated many different animal species, the first of which was probably the dog some 12,000 years ago. Felis silvestris was almost the last species to be tamed, being added to the list of domesticated animals after goats, sheep, cattle, pigs, chickens, horses, and even water buffalo. However, from these modest and comparatively recent beginnings, the cat is now on the verge of becoming the Western world's most popular pet; current predictions are that cats will soon overtake dogs as the most commonly kept pet. According to the Pet Food Institute in Washington, D.C., cats already outnumber dogs in the United States. In 1997 there were an estimated 70.2 million pet cats in the United States, compared with 55.9 million dogs.

Though many millions of cats are well-fed, well-loved family pets, millions more are feral, scavenging human leftovers. Many others live with and are fed by humans but supplement their diets with birds, rodents, and lizards.

Cats are believed to have been first domesticated in Egypt, some four thousand years ago, but the problem of identifying the exact period when domestication occurred is complicated by the fact that domestic cats are only recently descended from the African wildcat (Felis silvestris lybica). For this reason, the skeletons of domestic cats and wildcats are difficult to differentiate. When cat bones are excavated from archaeological sites, it is difficult to establish whether they belong to wildcats that were scavenging and hunting around human dwellings or to domesticated wildcats living with people.

Though most of the evidence points to Egypt as the birthplace of the domestic cat, bones of small felids have been found at older archaeological sites in other areas. The remains of African wildcats have been excavated from Jericho and dated at 6000 to 7000 B.C., but there is no evidence that these were from domesticated animals; rather, they may have been the bones of cats killed for fur.

Recent excavations of a six-thousand-year-old settlement on the Mediterranean island of Cyprus have unearthed a cat's jawbone, suggesting that cats may have been associated with people for longer than was previously thought. Wildcats do not occur naturally on Cyprus, so the animal on the island must have been brought there in a boat, either as an accidental stowaway or as a pet.

Archaeological evidence shows that African wildcats were certainly spending time around Egyptian towns and villages some four to five thousand years ago, but their exact status and the process of domestication remain unclear. One theory is that wildcats simply began to hang around farms, granaries, and town middens, drawn by the abundant rodents that were attracted by the grain and garbage. Historical and biblical accounts record plagues of rats and mice decimating grain stores and spreading disease, and almost any predator that reduced the numbers of these rodents would undoubtedly have been encouraged.

Others argue that the most likely route to domestication was through people taming captured kittens, just as many South American and Asian people tame monkeys and birds today. It is known that the Egyptian people of that time had an extraordinary passion for taming wild animals, and it was common for wealthy families to have large menageries containing baboons, lions, mongooses, hyenas, and gazelles. Given that cats were objects of worship and thought to be the earthly representatives of various deities, it is highly likely that the Egyptians attempted to tame them in order to add them to their animal collections.

For the Egyptians, the process of domestication was aided by the fact that the local subspecies of wildcat, Felis s. lybica, was a much less aggressive animal than the virtually untamable European wildcat, F. s. silvestris. Even so, pure F. s. lybica kittens are reported to be quite difficult to handle. The veteran South African zoologist Reay Smithers kept several purebred F. s. lybica, along with some lybica-domestic crosses. Smithers wrote that "the progeny of Komani and a pure male from Botswana, Igola [both F. s. lybica], were long-legged with red ears and from the earliest stages, unhandleable, spitting and scratching or diving for cover when approached." However, Smithers adds that crosses between domestic cats and wildcats are easy to handle and tame easily. When Smithers's wild-caught female mated with a domestic cat, the offspring "turned out to be splendid house cats, great hunters and reached an adult weight of 12 to 14 pounds, some two pounds heavier than their mother."

Beginning about two thousand years B.C., the domestic cat's history becomes easier to follow, as this marks the time when Egyptian artists began to depict the cat in mosaics and paintings. Statues, amulets, and pictures show cats in a variety of contexts, sitting under chairs, riding in boats, and being worshipped as deities. In Thebes, in the tomb of Nakhte, dating to 1415 B.C., there is a painting of a cat killing a mouse under its owner's chair. In another tomb dated about 1900 B.C., the bones of seventeen cats were discovered, along with several small pots believed to be for offerings of milk.

At that time in Egypt, the cat was associated with a confusing array of gods and religious beliefs. One papyrus depicts the sun god Ra as a cat with spots and barred markings, holding a knife in its paws. In the drawing the cat is cutting off the head of the serpent of darkness, who was believed to swallow the sun every evening. This association with the sun apparently gave rise to the string game "cat's cradle," versions of which are played all over the world. Though the meanings of the various patterns have been obscured by time, the string cradles were used as a means to control the movements of the sun.

Lions were also associated with the sun god, and cats were seen as closely related to lions. The lion-headed goddess Sekhmet represented the destructive aspects of the sun and was associated with wrath and vengeance. According to one version of the myth, Bastet was the sister of Sekhmet, the daughter of Isis and Osiris the sun god. Though Bastet was originally lion-headed, she became more and more frequently depicted with a cat's head and came to represent the good and benevolent aspects of the sun. Bastet eventually became the great cat goddess who was in charge of all growing things; she was a symbol of fertility for both crops and women, and eventually came to be known as the goddess of joy and love.

Around 1000 B.C. Bastet emerged as a major deity and the focus of the famous cat cult. The center of the cult was the great temple of Bastet in the city of Bubastis, which is east of the Nile delta and is today marked by a mound called Tel Basta. Herodotus, who visited the city in 450 B.C., describes the shrine of the goddess Bastet as "standing on an island completely surrounded by water except at the entrance passage." According to Herodotus's detailed description, the shrine itself was built of fine red granite and encompassed a sacred enclosure about 600 feet square, beyond which was a larger enclosure containing a canal, a grove of trees, and a lake. In addition to a huge statue of the goddess Bastet, the shrine contained thousands of cats, which were fed and cared for by innumerable priests and attendants.

As the goddess of joy and love, Bastet was an extremely popular deity, and each year thousands of people celebrated the festival of the cat goddess with a pilgrimage to Bubastis. The event was one of the principal festivals in Egypt and seems to have been somewhat akin to a weeklong party; the mood was festive, and there was much drinking, singing, and dancing.

During the cat cult of Bastet, images of cats were carved and sculpted in every material from gold to mud. Paintings depicted cats of various colors, including ginger, orange-brown, and gray tabby. Cats were shown eating fish, springing at waterfowl, and catching mice. Bronze cat statues were used as votive offerings at shrines, and cat amulets made of gold, glass, jasper, and stone were worn around the neck and buried in cat graves. The penalty for killing a cat was death, and people would flee if they saw a sick or injured cat in the street for fear of being held responsible for the creature's demise. Cats were also highly esteemed as pets; many people owned cats, and the death of one of these pets sent the entire family into mourning. Behaving almost as if a human family member had died, people shaved their eyebrows as a sign of respect and had the dead animal embalmed and buried in a special cat cemetery. The embalming procedure and funeral trappings varied depending on how wealthy the family was. A poor man's cat was rolled in a piece of plain linen, whereas a rich man might commission an elaborately embalmed cat mummy with a decorated papier-mâché mask. Thus embalmed, the body was placed in a mummy case, or if it was a kitten, in a small bronze coffin. Food for the afterworld, in the form of mummified mice and small pots of milk, was buried with the cat.

It was during this period that the great cat cemeteries were laid out along the banks of the Nile, where huge underground vaults and repositories held the mummified remains of several hundred thousand cats. One such burial ground was discovered at Beni Hasan in 1888 when a farmer accidentally dug into a vault containing thousands of mummified cats. The contents of this particular vault were so numerous that a businessman hired people to strip cloth and dried fur from the bones so that the bodies could be turned into fertilizer. Nineteen tons of mummified cat bones, or the remains of some eighty thousand cats, were shipped to Manchester in England to be ground up for use as fertilizer.

Fortunately, a few of these cat skeletons survived to be examined and described by scientists. Eighty-nine skulls from Beni Hasan have been dated from 1000 to 2000 B.C., and of these, four or five are thought to belong to Felis chaus, the jungle cat, while the rest are Felis s. lybica. Another collection of skulls and mummified animal remains from Egypt, dating from 600 to 200 B.C., was presented to the British Museum in the early 1900s, but the box containing the specimens was put into storage, misplaced, and only rediscovered some fifty years later. When examined, the collection was found to contain one hundred and ninety-two cats, seven mongooses, three dogs, and a fox. Three of the skulls were from Felis chaus, the jungle cat, but the remainder were those of the African wildcat.

The Egyptians kept their cats under close guard, and by making their export illegal, essentially prevented the spread of domestic cats to other countries. The earliest record of a domestic cat in Greece is a 500 B.C. marble bas-relief scene of a cat on a leash confronting a dog. This must have been an unusual event, because at that time cats were almost unknown in Greece and Rome; ferrets were the animals of choice for rodent control. Cats remained rare until the fourth century A.D., when the Roman writer Palladius recommended cats as an alternative to ferrets for getting rid of moles in artichoke beds.

F. E. Zeuner, in his classic work A History of Domesticated Animals, suggests that cats began to spread from Egypt to the rest of the world soon after Christianity arrived in Egypt because this change released the restrictions on the movements of cats and eventually led to cats being imported to Rome. Others believe that the cat's spread through Europe was linked to the spread of the brown rat and the house mouse. Once the cat had arrived in Rome, it was almost inevitable that it would spread throughout Europe, quite likely as a camp follower and companion to the constantly traveling Roman armies. The domestic cat was introduced to Britain by the Romans, and the remains of cats have been found in many Roman settlements in England.

By the tenth century the cat was becoming more common throughout much of Europe. In Wales in the tenth century, a hamlet was defined as a place that contained "nine buildings, one herdsman, one plow, one kiln, one churn, one bull, one cock and one cat." According to the laws of Hywel Dda, a Welsh king who lived about A.D. 945, a cat was worth four pence. Thus it had the same value as a dog, but was worth more than a small pig, a lamb, or a goose, each of which were said to be worth one penny. In Germany in the twelfth century the punishment for killing another person's cat was a fine of sixty bushels of corn.

Cats most likely spread through Europe and around the world by way of barges and sailing ships, and there are many nautical terms and weather descriptions that make reference to cats. A light breeze that ripples the surface of the water is known as a cat's paw, and a cat scratching the leg of a table or chair was thought to foretell a storm. Many other words with nautical associations began with the word cat, such as cat-o'-nine-tails, catboat, catwalk, and cat rig. Carried across oceans or walking from village to village, cats gradually spread across the globe, and by the tenth century they had reached Japan, by way of China.

Cats seemed to attract more than their fair share of myths and superstitions. In Scotland and Japan, tortoiseshell cats were believed to be able to foretell storms. People in eastern Europe thought that evil spirits took possession of cats during thunderstorms and that lightning was produced by angels in an attempt to exorcise the spirits. In that part of the world, cats were pushed outside as soon as a storm began so that the lightning need not strike the house to reach the cat. In other places, such as Indonesia, cats were used as rainmakers. They were carried three times around a dry field, then dunked into a container of water.

The Middle Ages marked the beginning of three centuries of persecution of the cat, and by the fourteenth century cats were in serious trouble in Europe. Long associated with the moon, cats were now considered to be the familiars of witches and disciples of Satan. Witches were believed to have a unnatural nipple with which they suckled their cats, and several witches are said to have confessed to feeding their cats milk and blood. At the trial of one woman in Essex, evidence was given of a cat who would "suckle bloud of her upon her armes and other places of her body." Women, especially the old and ugly, became special targets of investigation, and many were tortured and persecuted for being witches; their ability to transform themselves into cats was accepted as fact, and taken as evidence that they were witches.

Witches were thought to be able to bring all kinds of misfortune upon people, and as the servant of the witch, the cat-familiar took an active role in spreading the havoc. Therefore, an unknown cat on the doorstep often precipitated a crisis of confidence as the household wondered whether the creature was a cat or a witch come to cast a spell on someone. Witches were thought to be able to transform themselves into cats to disguise their activities, and some believed that witches rode to their midnight meetings on the backs of giant cats. The first official trial for witchcraft took place in 1566, and as a result Agnes Waterhouse and her daughter Joan were executed as witches. It was said at the trial that the women had "a whyte spotted catte" and that they "feed the sayde cat with breade and milkye . . . and call it by the name of Sathan." Hundreds of years later cats were still associated with the devil and bad luck, and were often viewed as the witch's familiar or companion who carried out her evil plans. Even the well-known saying that a cat has nine lives has its origins in witchcraft. According to a book titled Beware of the Cat, which was written in 1584, "It was permitted to a witch to take on her catte's body nine times." Also connected with witchcraft is the old superstition that a black cat crossing a person's path brings bad luck; this originated with the belief that the black cat was marking a path to Satan.

It is astonishing that any cats managed to survive this period of European history. People killed cats whenever they encountered them, and one of the most common events of a feast day was the torture and death of any cat that allowed itself to be captured. The act was considered a symbolic way to drive out the devil. During this time cats were burned, beaten, drowned, and generally tortured and abused. For nearly three centuries the cruel treatment of cats was encouraged, as it was thought to be amusing and entertaining. One popular instrument of public torture, known as the cat organ, was said to have been invented in Brussels in 1549 for a festival in honor of Philip II. The device consisted of twenty cats confined in narrow cases with cords attached to their tails. The cords were attached to the keyboard of an organ so that when a trained bear pounded on the keys, the cords jerked the tails of the cats and made them howl. Another unpleasant piñata-like contest survived in Scotland until as recently as the end of the eighteenth century. During this game a cat was placed in a hanging barrel of soot, and men on horseback took turns riding under the barrel, striking at it until a hole appeared and the cat leaped out and ran away.

Cats did not return to public favor until the seventeenth century, when the French Cardinal Richelieu took to keeping dozens of them at court. Other people followed suit, and the cat soon regained some of its former popularity in Europe. However, despite the fact that today cats are on the verge of surpassing dogs as the world's most popular pet, many people still dislike them. Surveys of attitudes in the United States show that there are nearly seven times as many cat haters as there are dog haters, and many people associate cats with allergies.

One of the earliest descriptions of human allergy to cat hair comes from The history of four-footed beasts, written by Edward Topsell in 1607. Topsell writes that "the breath and favour of Cats consume the radical humour and destroy the lungs, and therefore they which keep their Cats with them in their beds have the air corrupted, and fall into several Hecticks and Consumptions."

Both the Egyptian and Chinese words for "cat" seem to have been derived from the animal's vocalization; in Egypt the cat was called mau, and in China it was mao. Other names for the cat have been derived from the Nubian word kadiz, from which we get the English word cat, the French chat, the Greek kata, the German katze, Spanish gato, Italian gatto, Russian koshka, and Arabic qutta. Other words and names for the cat derive from Pasht, which comes from Bastet; these include the English puss and the Romanian pisicca.

Description

Today's domestic cats have a much greater variety of coat colors, textures, and patterns than they did two hundred years ago, but the majority of the world's cats still have some variation of the tabby coat pattern. The tabby pattern is closest to the wild, or agouti, pattern, and on close inspection each hair can be seen to be made up of several different colors, including brown, black, gray, or white. The coat color of the domestic cat is thought to have originated by mutation from the original striped tabby coat of the African wildcat.

The selective breeding of cats began comparatively recently, in the late nineteenth century. The move to breed new types of cats began in Britain and coincided with a surge of public interest in the newly introduced theory of evolution and a widespread fascination with the idea of improving and enhancing different breeds of animals.

Modern cat breeds originated from two main types: the European, or cold-weather, type, which has a large head, stocky body, and thick coat and resembles the European wildcat, and the Foreign, a warm-climate type with a slender body, long limbs, large ears, and short coat that resembles the African wildcat. There is no exact definition of what constitutes a "breed," but in general, body conformation, geographic origin, color, and hair type are the major criteria used to define the characteristics of a breed. Many of today's breeds are just varieties of existing breeds that have been given fancy names: the Balinese is really a long-haired Siamese, and the Somali is a long-haired Abyssinian. There are many other unusual-looking breeds of domestic cats, including the tailless Manx and the Japanese bobtail, both of which have existed for many centuries. More recent breeds with characteristic physical features include the Scottish Fold, with lop ears, the Cornish Rex, distinguished by its short, curly coat, and the Sphinx, a hairless cat. With only about a hundred years of selective breeding behind them, domestic cat breeds are just beginning their evolution, and it will be some time before they acquire the variety and dramatic differences that today's domestic dogs have.

Folklore has long maintained that cats always land on their feet, and a study by the Animal Medicine Center in New York has now confirmed that cats do indeed manage to right themselves during a fall from a building or ledge. Doctors in the New York veterinary hospital looked at 132 cats that fell from tall buildings, from the second to the thirty-second floor. Astonishingly, 90 percent of the cats that fell from these heights survived, and almost two-thirds of them required no medical treatment. By comparison, people falling more than six stories are almost invariably killed. Cats survive these incredible falls because they turn their legs downward and extend their limbs outward, essentially assuming a flying squirrel or gliding position. This prevents them from tumbling head over heels through the air while falling and saves them from hitting the ground headfirst. The legs-out position also changes the aerodynamic drag on the cat's body and slows it down so that it hits the ground with the least possible force. One of the record falls involved a cat from New York named Sabrina, who fell from the thirty-second floor of an apartment building and landed on a concrete sidewalk. Sabrina walked away from the incident with a chipped tooth and a minor chest injuryan achievement not many other animals could match.

Distribution

From their beginnings in Egypt, the Middle East, and Europe, domestic cats have accompanied people to almost every corner of the globe. Wherever people have traveled, they have taken their cats with them. Geographic features such as major rivers and oceans that are barriers to most animals have the opposite effect on cats. Almost as soon as people began to move goods around on ships, cats joined ships' crews. These cats traveled the globe, joining and leaving ships at ports along the way. Because the coat colors of cats do not occur with equal frequency all over the world, geneticists were able to create distribution maps for the most common color morphs. The European distribution of the blotched tabby coat patterna relatively new mutation that has not yet spread worldwideshows that this type has its central focus in Britain and exists at high frequencies across central France and Germany. This pattern seems superficially at odds with the dispersal by water theory, until the routes of the rivers Seine and Rhone and their barge canal systems are overlaid on the distribution. There is also a suggestion that the distribution of the blotched tabby in New Zealand, Canada, Australia, and Tasmania reflects British migration. As settlers left to take up residence in their new countries, they took their placid-natured blotched tabbies with them. In addition to going along as traveling companions with colonists, cats also accompanied sailors on their voyages, with serious consequences for the native fauna wherever they disembarked. Time and time again the scientific literature tells the sad story of domestic cats let loose on remote islands, where they became serious predators on native wildlife. There are now major cat eradication programs in the Galápagos, New Zealand, Australia, and other places.

Ecology and Behavior

Domestic cats are opportunistic predators, the ultimate generalists. They can live indoors year-round in a small apartment or outdoors, supplementing their diet of domestic cat food with birds and rodents. Though cats have been domesticated for some time, they quickly and easily revert to the wild or feral state. In the last hundred years cats have spread to virtually every corner of the globe, and today, feral cats live on remote subantarctic islands, in forests and farmlands, in suburbs, and in industrialized cities.

Feeding Ecology

Domestic cats hunt mainly on the ground, but they can also climb well. Cats are famous for their stealthy approach to prey. In mythology cats are often associated with thieves because of their ability to creep around unnoticed; in Sanskrit, the same word is used for both cats and thieves. A cat setting out for a hunt often appears to know where it is going. When it arrives, it begins to search, moving more slowly, looking around or crisscrossing the hunting area. If it is hunting rodents, the cat will sit and stare at a hole; when a mouse appears, the cat will usually wait until the prey has moved some distance from its burrow before pouncing.

Hunting birds requires different tactics. The cat stalks as near as possible, slinking silently toward the victim, belly to the ground, then stopping. All the time the cat is a picture of concentration, ears pricked, body tensed, eyes fixed on the intended prey. As the cat gets closer, it gets more and more excited; the tip of its tail twitches, its bottom wiggles, and its hind feet tread the ground as if seeking firm purchase. With eyes still fixed on the prey, the cat sometimes moves its head from side to side. Folktales have it that the cat is trying to mesmerize its intended victim, but in reality it is only trying to gauge the distance as accurately as possible. The final rush is fast and determined, an all-or-nothing gambit. The hind legs provide the power for the leap, and the front legs are thrust out to grab the prey. If the pounce connects, the cat slams the prey to the ground with its forelegs and delivers a swift bite to the neck. Unless they are chasing grasshoppers or butterflies, cats rarely leap into the air to catch something; even when they are hunting birds, cats prefer to keep their hind legs planted firmly on the ground. This gives the cat a stable footing; if the prey starts to fight or is difficult to hold down, the cat can grapple without losing its balance.

Domestic cats are specialized predators, but they are versatile generalists in terms of what they eat. Where they are not subsisting on human-provided foods or refuse, cats take a wide variety of prey and readily switch prey with changes in availability. Excluding those on islands, the vast majority of cat studies show that small mammals, principally rodents, and rabbits or hares are the dominant prey. Adult rabbits are occasionally taken, but they appear to be near the upper limit of prey size; thus predation on lagomorphs is heavier on young of these species. Similarly, adult Norway rats are sometimes killed by cats, but laboratory studies show that few cats will attack an aggressive adult Norway rat. Despite frequent accusations, birds do not figure prominently in cat diets except on islands. Surprisingly, predation on reptiles, primarily lizards, appears to be important both on equatorial islands and on islands at much lower latitudes.

The effects of predation by domestic and feral cats on populations of small mammals are not well documented, but several studies suggest that it can be an important factor. Where Norway rat numbers had been reduced severely by human efforts, cat predation was able to hold the population at these low levels, but at high rat densities cat predation was ineffective as a means of control. Similarly, experiments with enclosures have shown that predation, especially by cats, can hold rabbit numbers below those set by the food supply. Predation by cats is also thought to be largely responsible for the 3- to 4-year cycle of vole numbers in California.

Predation by cats can have a serious impact on wildlife, especially on islands where the native fauna has evolved in isolation from predators or where seabirds nest on the ground. Today in the Galápagos Islands, native rodents exist only on those islands where there are no cats. Similarly, cats have eliminated several island species of birds, and others, such as the saddleback and New Zealand's kakapo, a large flightless parrot, are teetering on the brink of extinction largely due to predation by feral cats.

There are several well-documented cases of local extinction caused by cat predation on seabirds, especially on islands. At one time Ascension Island in the Atlantic Ocean was home to vast breeding colonies of several different species of seabirds, but once feral cats were introduced, the birds ceased to nest there. Remnant populations of ten other species of seabirds breed on isolated rocks around Ascension, but today only sooty terns breed on the island. In New Zealand feral cats eliminated large breeding populations of diving petrels and broad-billed prions from Herekopare Island. In 1949 five cats were introduced to Marion Island as pets of the members of a meteorological expedition; by 1975 an estimated 2,139 cats were thought to be killing about 450,000 burrowing petrels a year. Three of twelve species of petrels that breed on Marion Island were eliminated by cats in less than fifteen years.

Cats have also caused extinctions among island reptile populations. An estimated 15,000 Turks and Caicos Island iguanas were wiped out between 1974 and 1976 when a large hotel was built and domestic cats and dogs were introduced. On the Galápagos Islands the endemic marine iguana is endangered on several islands because of the combined effects of predation by introduced cats, dogs, rats, and pigs.

Cats can also be major predators of birds and small mammals in urban areas. In an amazingly detailed study of domestic cats in the small English village of Felversham, two scientists managed to persuade the owners of seventy cats to record and save all the prey items their pets brought home. These cats brought in a total of 1,090 prey items during the year-long study. There was great variation in individual hunting success: six cats never brought anything home; one cat brought in ninety-five items. Wood mice were the most frequent prey (17 percent), followed by house sparrows (16 percent), field voles (14 percent), and common shrews (12 percent). Five species made up 66 percent of the animals caught. The cats caught fewer prey in winter and on wet, windy days. At least 30 percent of sparrow deaths in the village were due to cats.

Though the idea is not popular with cat owners, there are indications that domestic cats can push small mammal populations to unnaturally low levels, which in turn may affect hawks and other birds of prey. Buffered by cat food and handouts from humans, cats can continue to hunt sparse rodent populations for longer than a wild predator could. Therefore cats fed by people could potentially reduce small mammal populations to a point where they would be unable to support wintering birds of prey.

Social Organization

Feral domestic cats are common and fairly accessible compared with other species of wildcats, and there have been several long-term studies of their social systems and population dynamics. Information on many aspects of the biology of domestic cats, including their social life, predatory behavior, reproduction, and the development of young, is discussed in the excellent volume edited by Turner and Bateson. Some of the best studies are those by Dards in a dockyard in southern England, Liberg in rural Sweden, Macdonald on English farms, Natoli in a city park in Rome, Izawa in a Japanese fishing village, Jones and Coman in the Australian bush, and Corbett on a remote island off the coast of Scotland. The types of prey available to cats in these studies varied from rabbits and small mammals to fish offal, garbage, and cat food left by visiting cat lovers.

Though their social organization is sometimes dismissed as a product of domestication, domestic cat society provides us with an excellent model of the way in which ecological factors shape social systems. Domestic cats live in habitats as varied as city apartments, oceanic islands, and barnyards. Their social systems must accommodate densities that may vary from one cat per square kilometer to more than 2,350 cats per square kilometer. Domestic cats may live solitary lives within a dispersed population of cats or in groups around a dumpster or other clumped food resource. As with other felids, females compete for food and safe den sites to rear young, while males compete for access to females. The result is a social system that is driven largely by the abundance and predictability of the food supply and, to some extent, the timing of female estrus.

Most studies have not measured food availability, but it is clear that the land tenure system of domestic cats varies with the distribution of food. Indeed, cat populations can be divided into those in which females form groups and those in which females live alone. Where the food supply is sparse but evenly distributed, females live in nonoverlapping territories. Males occupy larger ranges that overlap the ranges of several females. Where food is plentiful and clumped, such as at the offal dumps in a Japanese fishing village, females live in groups and males are solitary. Cats feeding solely on natural prey have not been found living in groups, presumably because natural prey is never clumped or abundant enough.

Home range sizes of domestic cats mirror food availability. Female range sizes may be as small as 0.1 hectare at a fish offal dump in Japan or as large as 270 hectares in the dry grassland of Australia. At these same sites, male ranges varied from 0.31 hectare in Japan to more than 900 hectares in Australia. In some regions female domestic cats are seasonal breeders, and in these populations breeding males expand their ranges during the mating season to gain access to as many females as possible. Breeding male home ranges are usually three to four times larger than female ranges, while subordinate male cats generally have smaller, female-sized ranges.

One of the most intriguing characteristics of domestic cat social organization is the fact that, like lions, these diminutive felids are able to live in social groups when conditions are right. Food seems to be the key factor favoring group living, and it is notable that all the documented incidences of domestic cats living in groups occur at clumped, abundant, artificial food sources such as dairy farms or garbage dumps.

In the early 1980s Olof Liberg conducted a now classic field study of the domestic cat's social behavior. In a rural area of southern Sweden, Liberg radio-tracked thirty-seven cats, which he classified as either feral, if they hunted for themselves and were not fed by humans, or domestic, if they were attached to and provisioned by households. Only three female cats were feral, and they had home ranges of about 200 hectares, about four times the size of the ranges of their domestic female counterparts. Feral females also spent about twice as much time hunting as domestic females.

Domestic females lived alone or in groups of up to eight related adults. These females shared a small communal home range of about 50 hectares, and there was little overlap between females of different kin groups. Females of different kin groups behaved aggressively toward one another and excluded nonkin from "their" household. The household represented a clumped and defensible resource that provided the kin group with essential resources, food, and shelter.

Among males, young pre-dispersal males had the smallest ranges and were the least active, while breeders had the largest. Domestic male breeders had ranges averaging 380 hectares, whereas a feral breeding male covered an area of 990 hectares. Most domestic males were subordinate to the local feral males. Male ranges overlapped considerably.

At almost the same time that Liberg was following his cats in southern Sweden, Macdonald and Apps were beginning a study of a colony of farm cats in England. Macdonald and his co-workers focused more on quantitative observations of social behavior, using scan sampling to record the activities of the cats. The three females in this group had ranges averaging 13.1 hectares, while the adult male's home range was 83 hectares. The females gave birth to kittens in communal dens, and all tended, groomed, and nursed one another's young. In this colony the most obvious benefit of group living seemed to be alloparenting behavior. However, the costs of this behavior were also high, as at least two litters succumbed to feline panleucopaenia, an infectious viral disease, which is well suited for transmission at communal dens.

As with lion prides, communal denning helps female domestic cats defend their young against infanticidal males, which is strongly suspected to be an important factor in group formation. Though female domestic cats may sometimes live like lionesses, tomcats do not form male coalitions and associate like male lions, probably because they do not need to. A single barnyard tomcat can roam over two or three colonies of females, and unlike the male lion, which often relies on the lionesses of the pride to capture prey, he can catch his own food.

Male lions and male domestic cats do share one behavior that is rare in the felid family. In both species, several males may take turns mating with a female in heat. Contrary to what one might expect, fights are rare, but an older dominant male probably gets most of the matings. With the lion, this male tolerance can be explained by the fact that male coalitions are almost always relatives, but this is not so for male domestic cats. It is still unclear why domestic cats and lions should behave similarly in this regard. Though the techniques are now available to investigate paternity, we do not yet know which male sires the most kittens, or whether the number and order of matings affects conception.

Where female groups exist and the relationships between group members are known, the social system is similar to that of a lion pride in that the group is made up of related females and is stable over the long term. Females in the group are unanimously hostile to outside females, but within the group they show distinct "friendships" or preferences for the company of other females. These affiliations appear to be governed by age, status, and blood relationships. Males usually leave or are driven out when they become sexually mature, and adult males are loosely attached to the female group. Several adult male ranges may overlap the range of the group of females, and in places where the food resource is superabundant, several groups of females with overlapping males may form a colony.

When it comes to breeding and raising kittens, females in a group may choose one of two strategies. Most rear their kittens alone, but there are cases of as many as four mothers pooling their litters and raising them together. Like lions, females in a group sometimes show a high degree of synchrony of estrus, which makes communal care more feasible, but in one recorded case, litters were seven weeks apart, meaning that the first litter was highly mobile and almost weaned when the last was born. Females who raised their young together were closely related.

Just as with lions, when female domestic cats cooperate to care for several litters, the behavior is thought to be an adaptation for defense against infanticidal males. Indeed, on the few occasions when it has been observed, infanticide in domestic cats has most often involved kittens from litters being raised by a solitary mother.

The social organization of domestic cats, like that of other felids, appears to rely on a system of visual, vocal, and olfactory signals. Each is used alone or in concert with others depending on the circumstances. Some studies suggest that cats avoid using common areas at the same time based on scent marks, but among cats living in the same colony, there was little evidence that scent functioned to keep animals apart. Adult male cats spray urine frequently while traveling, leaving their marks at locations where other cats are likely to encounter the signals. In Sweden, dominant males sprayed at rates of 22 marks per hour, compared with 12.9 urinations per hour for subdominant males. Feces deposited within the core areas of male ranges were always covered, but outside of those areas they were frequently left uncovered. Similarly, female farm cats were more likely to cover their feces within the core of their ranges. Females also spray urine while traveling, but not at the high rates recorded for males. Females spray at higher rates just prior to estrus, thus ensuring a male's presence at the appropriate time.

Scent marking in cats also includes rubbing the cheeks, chin, and flanks on objects, on urine marks, and on other cats, a behavior that facilitates leaving one's scent and being marked with the odor of other cats as well.

Cats also communicate by vocalizations, and their vocal repertoire includes about a dozen calls. Most calls function in close-range situations, including the purr, hiss, spit, growl, meow, and gurgle. The yowl or "caterwaul" is most often heard in the context of sexual activity. Body and tail postures, as well as facial expressions, are also used as signals in close-range encounters. Leyhausen has described many of the postures and expressions associated with agonistic behaviors in cats.

Reproduction and Development

Female domestic cats are seasonally polyestrous, and the onset of sexual activity appears to be controlled by photoperiod. The main breeding season is in early spring at northern latitudes, in mid-January in Italy and in February-March in Sweden. During this period females typically have a series of estrous cycles at fifteen-day intervals unless conception occurs. Estrus, or the period of receptivity, lasts one to four days, during which a female may attract as many as twenty males. Females may be inseminated by as many as ten different males, and some litters have more than one father. In most areas domestic cats commonly have two litters a year. In Sweden, for example, 60 percent of older females bred twice a year, and 7 percent bred three times.

Toward the end of the sixty-three-day gestation period, the female begins to search for a safe place to deliver her kittens. Cats are highly individual in their choice of birth dens. A cat owner may provide the expectant mother with a secluded box, only to have the cat choose the back of a closet or the center of the bed. Feral cats make their dens in thickets, rockpiles, or almost any safe, out-of-the-way site. As labor begins, the cat becomes restless, licking her vulva and changing positions. Delivery times are highly variable; it may take as long as fifty minutes or as little as one minute for a kitten to be born, and the interval between successive births in a litter can be similarly variable.

Kittens are born wrapped in a transparent membrane, which the mother removes and eats. She licks the kitten's nose and mouth to clear any mucus that might impede breathing, then licks the kitten dry. Because it is likely that a mother cat may be attending to a littermate and not attend to a kitten immediately after it is delivered, newborn kittens have the ability to survive without oxygen within the birth sac for a short while after birth. Some scientists have suggested that the newborn kitten's tolerance for lack of oxygen is the reason why kittens are reputed to be difficult to drown. After the whole litter is born, the female usually eats the placenta, thus cleaning the nest of anything that might attract predators.

A female cat can give birth to anywhere from one to ten kittens, but an average litter consists of four or five. The largest litter ever recorded consisted of thirteen kittens. The young weigh about 90 to 110 grams at birth, or roughly 3 percent of the mother's body weight. In general, individual weight declines as litter size increases.

Kittens are born with closed eyes, small, flattened ears, and a poorly developed sense of hearing. For the first two weeks, life is dominated by the senses of smell, touch, and temperature. Within an hour of birth each kitten finds a nipple and begins to nurse. By the second or third day members of the litter have established teat ownership, and each kitten suckles exclusively from its own nipple. This saves time, avoids fighting, and ensures there is a functioning nipple for each kitten. Suckling keeps the milk flowing, and within a few days the kittens have developed the characteristic "milk tread" in which they knead their mother's belly with their front feet. This motion stimulates milk flow and remains a part of the cat's behavior for life. Some older cats retain this kneading behavior to such an extent that anything soft and warm will set them off, and they seem unable to settle down to sleep without a few minutes of kneading. Others cats rarely seem to tread.

For the first few days of life kittens may spend as much as eight hours a day suckling; nursing bouts last as long as forty-five minutes, and the mother does not usually leave the nest for the first forty-eight hours. She keeps the kittens warm and licks them to clean them and stimulate defecation. For the first three weeks after birth the mother initiates suckling by purring and lying down in a characteristic nursing position in which her nipples are easy to reach. Later, when the kittens become more mobile and the weaning process begins, the kittens initiate suckling as they run to their mother when she returns to the nest. The first kitten to begin nursing also begins to purr loudly, and the rest of the litter comes running to join in. Griff Ewer called this the "dinner gong" function of purring.

Most kittens open their eyes sometime during their second week of life, and vision begins to play a major role in their lives when they are about three weeks old. A kitten's eyesight gradually improves until the fluids in the eye clear at about five weeks, and visual acuity continues to improve until the cat is three or four months old. There does seem to be considerable natural variation in the time when kittens first open their eyes, and the range is two to sixteen days after birth. Experiments in the laboratory have shown that several factors influence this timing, including paternity, exposure to light, the kitten's sex, and the age of the mother. Kittens reared in the dark opened their eyes earlier, as did kittens of younger mothers and female kittens; however, paternity was the most important factor, suggesting that the timing of eye opening is strongly influenced by genetics.

The milk of domestic cats contains eight times more protein than human milk and three times as much fat. Kittens grow rapidly as their mother uses her body reserves to produce milk. One study showed that mothers lost an average of 5.7 grams per day when lactating. As might be expected, kittens from larger litters are smaller and put on weight more slowly than kittens from smaller litters. Kittens in litters of seven or eight had a mean growth rate of 7.3 grams per day, whereas the mean growth rate nearly doubled to 13.7 grams per day for kittens in litters of two. In general, kittens double their birth weight in a week, triple it in two weeks, and quadruple it in three weeks. Initially, males and females grow at the same rate, but males start to grow faster at about eight weeks of age.

By the time they are three weeks old, most kittens are beginning to acquire the ability to regulate their body temperature, and this allows the mother to spend more time away from the birth den. At seven weeks of age the kittens are able to thermoregulate as well as an adult. Feral domestic cats begin to bring prey back to their kittens from the fourth week onward, and by the time they are five weeks old, the kittens start to kill mice. Most kittens are introduced to solid food at four to five weeks of age, and this event generally marks the beginning of the weaning period. Most kittens are weaned by the time they are seven weeks old, but may continue to suckle for several more months, particularly if the litter is small. As with most other features of the cat's life, the timing of weaning is highly individual. Some kittens continue to initiate nursing bouts long after they are physically weaned.

Kittens are fairly inactive for the first two weeks of life and usually begin to walk during the third week. Teeth begin to emerge shortly before two weeks of age and continue until the fifth week. A kitten's adult teeth begin to appear when it is about three and a half months old. By week five the kittens can run, and at eleven weeks they can complete complex tasks such as walking down a narrow branch and turning around when they get to the end. Kittens begin to play when they are about four weeks old. At first they play with littermates; later, when eye-paw coordination improves at about seven weeks, they begin to play with objects. The increase in object play coincides with the end of the weaning period, and if kittens are weaned early, the frequencies of certain types of play are increased.

It is commonly believed that play is an essential part of developing and honing future predatory skills, but experiments have shown that this is not necessarily true. Cats become competent predators through a variety of different experiences, and many different factors contribute to the development of predatory skills. Some kittens take a long time to become good at catching prey, and others are skilled predators from an early age, but these individual differences do not generally continue throughout life, and kittens that are poor predators have generally caught up by the time they become adults. Experiments have shown that predatory skills can be improved by early experience, and for a kitten to learn to kill prey, the experience does not necessarily have to be "hands on." It seems that simply watching the mother or another cat kill a rat is enough to teach a kitten how to deal with live prey. This observational learning is facilitated if the cat performing the act is familiar to the observer cat. When dealing with live prey, kittens tend to follow their mother's selection, and willingness to try new foods is also strongly influenced by the mother. One scientist trained mother cats to eat bananas and found that when their kittens were offered a choice between a meal of familiar meat pellets and a meal of the unfamiliar banana, most of the kittens imitated their mother and ate the banana.

Under natural circumstances kittens form strong social bonds with close kin and familiar individuals, usually their mother and littermates. Domestic cats have incorporated humans into the social group and react to familiar people with affection, but cats vary considerably in their friendliness. Some are adventurous and sociable with strangers; others are timid and shy, running from any loud noise. The difference can be apparent even in two kittens from the same litter.

While some of the differences among cats in friendliness can be accounted for by individual variation, several studies have examined the effect of early handling. These studies looked at variables such as the age of the kitten when first handled, the amount of time a kitten was handled each day, how many different people were involved, and whether the routine involved feeding. In general, kittens that were handled during the "sensitive period" (which is from two to seven weeks of age) approached people more readily and were willing to be held for longer. The timing of this sensitive period for socialization would seem to pose a problem for cat owners, as most kittens are not adopted as pets until they are eight to ten weeks old, and indeed, many places have regulations that prohibit the sale of kittens younger than eight weeks. Furthermore, all the evidence suggests that kittens should not be taken away from their mothers until they are naturally weaned, at about eight weeks. Separating a kitten from its mother earlier can cause stress and behavioral problems. Studies have found, however, that as long as kittens are handled during the sensitive period, they will relate well to the people who eventually adopt them. Cats that were never handled at all as kittens can be tamed, but it is a very time-consuming task. Individual idiosyncrasies aside, the best family pets are most likely to come from litters that have been handled gently and played with from an early age.

While attempting to find out what makes one cat friendly and another timid, one study turned up an unexpected variable: Kittens fathered by some males were more friendly than others. Thus, friendliness in cats may be inherited.

Female cats become sexually mature between seven and twelve months of age. Domestic cats have a high reproductive potential. According to the Guinness Book of World Records, the most prolific domestic cat produced 420 kittens in her lifetime.

Domestic cats live longer if they stay indoors and are well looked after. One survey found a number of cats that were nineteen to twenty-seven years old. The oldest recorded cat in the Guinness Book of World Records was a female tabby who lived to be thirty-four years old.